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Ammi Visnaga (L.) Lamarck (Apiaceae): Associated Beneficial Insects and Implications for Biological Control, with Emphasis on the Bell-Pepper Agroecosystem

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Ammi Visnaga (L.) Lamarck (Apiaceae): Associated Beneficial Insects and Implications for Biological Control, with Emphasis on the Bell-Pepper Agroecosystem Biological Agriculture and Horticulture, 1989, Vol 6, pp. 24 1-268 0 144-8765/89 $10 0 1989 A B Academic Publishers Printed in Great Britain Ammi visnaga (L.) Lamarck (Apiaceae): Associated Beneficial Insects and Implications for Biological Control, with Emphasis on the Bell-Pepper Agroecosystem Robert L. Buggl" and L. Theodore Wilson2 'Division of Biological Control, University of California, 1050 San Pablo Avenue, Albany, CA 94706, U.S.A.; and 2Department of Entomology, University of California, Davis, CA 9561 6, U.S.A. ABSTRACT Toothpick ammi (Ammi visnaga (L.) Lamarck, Apiaceae), a summer annual of Eurasian origin, commonly occurs along agricultural field margins in the Sacramento Valley of northern California, typically flowering from June to August. Observations during 1979-1984 yielded 65 distinguishable insect taxa attending the flowers, including 60 groups that were at least in part entomophagous. Peak attendances occurred for Braconidae, Ichneumonidae, Sarcophagidae, Sphecidae, Tachinidae, and Vespidae during early morning hours. Hay alfalfa (Medicago sativa L., Fabaceae) was grown in replicated plots with borders ofeither: (1) toothpick ammi, (2)common knotweed (Polygonumaviculare L., Polygonaceae), (3) prostrate pigweed (Amaranthus graecizans L.. Amaranthaceae), or (4) Clean-cultivated edges. Borders of flowering toothpick ammi attracted numerous entomophaga, but had no effect on densities of minute pirate bug (Orius trisricolor [White], Hemiptera: Anthocoridae) occurring in alfalfa. Pepper (Capsicum annuum L., Solanaceae) was grown with borders of either: (I) Vegetative Ammi, from which the flowers were repeatedly clipped, (2) Normally-flowering Amwi; or (3) Clean-cultivated borders (control) to assess impact on abundance of entomophaga. Predator densities and efficiencies within the plots were assessed weekly by: (I) Weekly whole-plant visual inspection; (2) Use of sentinel egg masses of beet armyworm (Spodoptera exigua (Hiibner), Lepidoptera: Noctuidae) or omnivorous leafroller (Platynota srultana Walshingham, Lepidoptera: Tortricidae), glued to the apical foliage and later inspected for occupation, or damage, by predators; and (3) Chunks of tuna placed on cards at ground level and later inspected for occupancy by bigeyed bugs (Geocoris spp., Hemiptera: Lygaeidae). Whole-plant visual inspection revealed a complex of generalist predators; those found on sentinel egg masses were in decreasing order of frequency Orius rristicolor, Geocoris pallens StAl, Lygus spp. (Hemiptera: Miridae), G. africolor Montandon, Collops vitratus (Say) (Coleoptera: Melyridae), and Nabis sp. *Present address: University of Georgia, Department of Entomology, Coastal Plain Experiment Station, Tifton, GA 31793-0748, U.S.A. 24 1 242 ROBERT L. BUGG AND L. THEODORE WILSON (Hemiptera: Nabidae). Weekly inspection of whole pepper plants failed to indicate differences in Orius density due to border regime. Likewise, there were no significant differences among weed regimes in numbers of Orius per sentinel egg mass placed on pepper plants amid the plots. However, another trial indicated that Orius recruitment to egg masses was higher on pepper plants with immediately-adjacent Ammi. Per-plant abundance of Geocoris and predation efficiency (Geocoris per lepidopterous egg mass) were separately regressed against Geocoris per tuna bait. Statistically-significant relationships were obtained when data were assessed across all dates, employing the mean counts for each date. Pending further testing, toothpick ammi should be tolerated in field- and road-side settings, as it is seldom a problem in crops and apparently supplies sustenance to many entomophagous insects. I NTR ODUCTIO N Flowering weeds have frequently been cited as important food resources for entomophagous insects (see reviews by: Altieri & Whitcomb, 1979; Altieri & Letourneau, 1982; and Bugg et al., 1987). Many of the relevant studies have emphasized the value of flowering Apiaceae that supply dietary pollen and nectar, leading to increased longevity and fecundity in adult parasitic insects, Generalist predators may also rely to varying extents on nectar and pollen (Bugg et al., 1987). Toothpick ammi (Ammi visnaga [L.] Lamarck, Apiaceae), a summer annual of Eurasian origin (Munz & Keck, 1973) commonly occurs along agricultural field margins in the Sacramento Valley of northern California. Typically flowering from mid-June through mid-August, the plant features umbels of 5.0 to 11.4cm diameter, comprising numerous small flowers. We were particularly interested in the possibility of enhancing the density and efficiency of minute pirate bug (Orius tristicolor [White], Hemiptera: Anthocoridae), a generalist predator that attacks many pests of vegetable crops, and which can occur on toothpick ammi at densities as high as 22 per umbel (W. Barclay, unpublished data). Several Anthocoridae, including 0. tristicolor, have been shown to respond favorably to vegetational diversifica- tion (Barney et al., 1984; Letourneau & Altieri, 1983; Solomon, 1980), although this has not consistently been the case (Bugg et a/., 1987). Salas- Aguilar (1976) suggested the use of flowering plants to enhance biological control by 0. tristicolor, which is known to feed on thrips and pollen (Salas- Aguilar & Ehler, 1977). The present study was designed to assess patterns of visitation by insects to flowers of Ammi, and implications for biological control of insect pests in agroecosystems. FLOWERING AMMl VISNAGA AND BIOCONTROL 243 METHODS The present study comprises four sets of experiments or observations conducted from 1981-1984 in Yolo County, California. The first involved cataloguing insects attending flowers of Ammi. The second set concerned the pattern of daylight attendance by insects. The third involved hay alfalfa (Medicago sativa L., Fabaceae) grown in replicated plots with borders of Ammi or other weeds, and assessed seasonal patterns of floral visitation by entomophagous insects. Also, sweepnet samples were made in the alfalfa, to assess possible effects of weedy borders on densities of Orius tristicolor. The final set involved bell pepper (Capsicum annuum L., Solanaceae) grown in replicated plots with clean-cultivated margins or borders' of flowering or vegetative Ammi. We conducted a census of flower visitors to detect seasonal patterns of attendance, and detailed assessment of generalist predators occurring amid pepper foliage and at ground level. The latter data not only provided information on possible effects of flowering Ammi, but also contributed new information about generalist predators in the bell-pepper agroecosystem. List of Flower Visitors A list of taxa whose members visit Ammi flowers was compiled during the years 1979-1984, based on observations and collections made in Yolo County, California. Diel Patterns of Flower Visitation To investigate timing of flower visitation, on August 5th, 1981, insects were collected at half-hour intervals from dawn until after dusk (approximately 0600-1830 Pacific Daylight Time) from a specimen of Ammi discovered some 5 km east of Woodland, Yolo Co., growing in a 74-ha field of overwintered sugarbeet (Beta vulgaris L. convar. crassa [Alefeld] J. Helm, Chenopodiaceae). The toothpick ammi plant had 27 umbels, and was located near the northwest corner of the field, with a field of overwintered alfalfa 50 m to the west, and a stockyard 30 m to the north. At each half-hour interval, we collected for 15 minutes, using an aerial net (net aperture'diameter 38-cm, fine mesh) to try to capture all insects visiting the Ammi plant. Insects were preserved in ethanol, then sorted to taxa. 244 ROBERT L. BUGG AND L. THEODORE WILSON Alfalfa Experiments Several of the entomophaga observed visiting flowers of Ammi are also common in the hay-alfalfa agroecosystem. We therefore tested the possible role of toothpick ammi in attracting and sustaining elevated densities of these entomophaga. During 1984, plots of overwintered hay alfalfa (cv ‘Amador’ were used for further experiments (see: Bugg et al., 1987, for details on establishment practices). The experiment employed a 4 X 4 Latin square design, with 6.1 X 6.1 m plots of alfalfa separated by 6.1 m clean-cultivated alleys. East and west borders of the plots featured strips of 0.61 m width devoted to alternative regimes: (1) toothpick ammi, (2) common knotweed (Polygonum aviculare L., Polygonaceae), (3) prostrate pigweed (Amaranthus graecizans L., Amaranthaceae), and (4) pure alfalfa (control). The alfalfa was mowed only once, in mid April, during the course of the 1984 studies. Flood irrigation was performed as needed. Common knotweed and prostrate pigweed were established by allowing pre-existing weed borders to reseed naturally, and later uprooting unwanted plant species. Ammi was established by two techniques. Ammi was seeded during mid-June, 1983, into seedling trays with 2.54 cm diameter cells, using a soil mixture of 3: 2: 1, sandy loam to delta peat to sand. Plants were raised in a lath house, watered as necessary, and transplanted to the appropriate plot borders during early July. Plants were installed, seven to a plot, at ca. 1 m intervals. At the same time, the same borders were sown with Ammi seed, approximately 50 g per border, which was incorporated by raking. The transplants provided flowers during spring of 1984, whereas plants that grew from seed provided flowers during the summer. Once flowering began, surveys of flower visitors were made during late- morning hours on sixteen occasions between days 108 and 228,
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